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Doctrine of Signatures: An Explanation of Medicinal Plant Discovery or

Dissemination of Knowledge?

Article  in  Economic Botany · September 2007

DOI: 10.1663/0013-0001(2007)61[246:DOSAEO]2.0.CO;2

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Doctrine of Signatures: An Explanation of Medicinal Plant
Discovery or Dissemination of Knowledge?'
BRADLEY C. BENNETT

Department of Biological Sciences and Center for Ethnobiology and Natural Products,
Florida International University, Miami, FL 33199

DOCTRINE OF SIGNATURES: AN EXPLANATION OF MEDICINAL PLANT DISCOVERY OR DISSEMINATION OF

KNOWLEDGE? The Doctrine of Signatures (DOS) is found throughout the world. Most
scholars dismiss it as a "primitive" or "prescientific" idea. Despite its long history, the doc-
trine has had little critical review. A careful evaluation of signatures suggests four things.
(1) There is no evidence that morphological plant signatures ever led to the discovery of
medicinal properties. Considering DOS in this manner is unproductive and largely untestable.
(2) Signatures are post hoc attributions rather than a priori clues to the utility of medicinal
plants. (3) It is productive to redefine signatures to include organoleptic properties associ-
ated with therapeutic value. Plants with strong odors or bitter tastes, for example, com-
monly are found in pharmacopoeias. (4) DOS should be considered for what it primarily
is—a way of disseminating information. DOS fundamentally is a mnemonic and, therefore,
is exceedingly valuable in traditional cultures.
Key Words: Doctrine of signatures, medicinal plant selection, medicinal plants, mnemonic,
organoleptic properties, transmission of knowledge, Paracelsus, signatures.

The Doctrine of Signatures (DOS) is a device used to transfer information, especially in

much–maligned theory that purportedly explains
how humans discovered medicinal uses of some
plants. DOS could be stated as form recapitulates
function—physical characteristics of plants reveal
their therapeutic value. For example, bloodroot
(see Table 1 for binomials, families, and signa-
tures of all cited taxa) contains the orange–red
alkaloid sanguinarine (Bennett et al. 1990). Ac-
cording to DOS proponents, the reddish color
indicates the plant's efficacy in treating blood ail-
ments. DOS also has a more specific meaning.
Some believe that the physical clues to a plants
use were given by the Creator.
The wild woods were full of creatures whose value
was written on each of them in the language of sig-
natures, if the seeker for simples could only manage
to decipher the label with which it had been consid-
erately tagged at the creation. (Eggleston 1901)
While DOS is often dismissed as primitive su-
perstition, I argue that it is primarily a symbolic
1 Received 18 October 2006; accepted 27 April 2007
preliterate societies. Shepard (2002) suggested a
similar idea when he wrote, "I hypothesize that
sensory evaluation serves as a mnemonic purpose
as well." Historical discussions of DOS are post
hoc interpretations by scholars who attempt
to explain why a plant is used for a particular
ailment. There is little evidence that a priori
analyses of morphological signatures led to
the discovery of any medicinal plant. Moreover,
this application of DOS is largely untestable-
researchers were not present when the utility of
medicinal plants was discovered. However, physi-
cal properties correlated with phytochemical at-
tributes (e.g., strong odors and the presence of
monoterpenes, bitter taste and the presence of al-
kaloids) may have led to the discovery of some
healing plants.
The Doctrine of Signatures is ubiquitous
(Balee 1994). According to Powell (1901), "All
American tribes entertain a profound belief in the
doctrine of signatures." Read (1927) wrote that
DOS is "to be found extensively quoted in the
Orient and Occident." DOS is alluded to in clas-
sical Greek literature on medicinal plants. Milton

Economic Botany, 61(3), 2007, pp. 246-255.

© 2007, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
 O
J

TABLE 1. EXAMPLES OF THE DOCTRINE OF SIGNATURES (BENNETT ET AL. 2002; CARLSON 1986; COLE 2004 [ORIG. 1656]; CULPEPER 1652; DELLA PORTA
1658 [ORIG. 1558] ; DIOSCORIDES 2000 [ORIG. A.D. 651 ; GERARD 1975 [ORIG. 1597]; ToUW 1982; GRIEvE 1971 [oRlc. 19311; MOONEY 1992 [ORIG. 1900];
READ 1927; ROWLINGS 1999; SAINE 2002; TYLER 1993). FAMILY DESIGNATIONS FOLLOW ANGIOSPERM PHYLOGENY GROUP (APG). N.A. = NOT APPLICABLE.

Common name Species Family Signature Medical Use

aconite Aconitum nape/lies L. Ranunculaceae eye like fruit ocular problem

akapmas Fittonia albivenis (Lindl. ex Deitch) Brummitt Acanthaceae liver—shaped leaves liver pain
bloodroot Sanguinaria canadensis L. Papaveraceae red—orange latex blood Fri
bugloss Anchusa officinalis L. (perhaps) Boraginaceae
z
seeds shaped like a viper's head venomous bites z
eyebright Euphrasia officinalis L. Orobanchaceae striped petals eye problems
foxglove Digitalis lanata Ehrhart, Digitalis purpurea L. Plantaginaceae heart—shaped leaves(?) cardiac problems H
ginseng Panax quinquefolium L. Araliaceae human—shaped rhizome panacea
herb scorpius Scorpiurus sp. (perhaps) Fabaceae coiled shoots resemble a scorpion's tail scorpion bites
O
machancha Columnea tessmannii Mansf. Geseneriaceae red—tipped leaves menstrual bleeding, snakebite H
mandrake Mandragora officinalis L. Solanaceae human—shaped roots panacea zrn
orchanet Alkanna tinctoria Tausch Boraginaceae viper—shaped seeds snakebite
puca panga Columnea ericae Mansf. Gesneriaceae red—tipped leaf menstrual bleeding, snakebite 0
purslane, common Portulaca oleracea L. Portulaccaceae stems worm—like anti—parasitic
saxifrage Saxifiaga spp. Saxifragaceae breaks rocks as it grows kidney stones G1
snakeroot Rauvoia serpentina (L.) Benth. ex Kurz Apocynaceae roots coiled, snake—like snake bite
snakewort Aristolochia serpentaria L. Aristolochiaceae leaf shape is similar to a snake's head snake bite C
soroci Momordica charantia L. Cucurbitaceae red aril blood tonic
stonecrop Sedum acre L. Crassulaceae leaf shape kidney stones
walnut, English Juglans regia L. Juglandaceae convoluted cotyledons brain ailments
wild comfrey Cynoglossum virginianum L. Boraginaceae fruits cling to skin memory
willow Salix spp. Salicaceae supple branches rheumatic joints
not cited Columnea crassa C.V. Morton Gesneriaceae red spots on leaves nose bleeding
not cited Magnolia spp. Magnoliaceae heart—shaped leaves cardiotonic
not c i ted Melissa officinalis L. Lamiaceae heart—shaped leaves heart weakness

N
rL^
248 ECONOMIC BOTANY [VOL. 61
(2002 [orig. 1667]) described the Archangel
Michael's use of eyebright to restore Adam's vi-
sion. Eyebright's distinctively characteristic striped
petals are supposedly reminiscent of bloodshot
eyes (Tyler 1993). DOS appears in current litera-
ture, as well. In Harry Potter and the Chamber of
Secrets, Madam Pomfrey revives petrified students
with a mandrake root decoction (Rowlings 1999).
Mandrake's signature is its human—like roots
(Mabberley 1987).
History of DOS through
the 19th Century
Pliny's Natural History contains the first writ-
ten record of the "pseudoscientific" Doctrine of
Signatures (Encyclopa:adia Britannica 2006a).
Dioscorides wrote, "... the Herb Scorpius resem-
bles the tail of the Scorpion, and [emphasis
mine] is good against his biting" (Dioscorides
2000 [orig. A.D. 65]). The phrase "and is good
against his biting" is very different from "and
therefore is used against his biting. Assuming the
latter is the common logical fallacy of post hoc
ergo propter hoc ("after this, therefore because of
this"), which is common in interpretations of
signatures.
Maycock (1991) claimed, "Galen was resolute
in his confidence of an underlying design
throughout nature to be demonstrable in accord
with a doctrine of signatures." Evidence for
Galen's adherence to DOS is weak. He is known
best for the concept contraria contrariis curan-
tur ("opposite cures the opposite"), the founda-
tion of allopathic medicine (Cooper 2004).
Paracelsus, the greatest DOS proponent, and his
followers espoused the position similia similibus
curantur ("like cures like"). He believed that
God provided signs within plants to indicate
their uses: "The soul does not perceive the exter-
nal or internal physical construction of herbs
and roots, but it intuitively perceives their pow-
ers and virtues, and recognizes at once their Sig-
natum" (Boericke 2004; Clarke 1923; Keightley
1925).
Bohme (1621) reiterated the Paracelsian the-
ory: "The greatest understanding lies in the sig-
natures, wherein man may not only learn to know
himself, but also the essence of all essences."
Nicholas Culpeper (1652) wrote, "... by the
icon or image of every herb, man first found
out their virtues." The 17th century botanist
William Cole (2004 [orig. 1656]) believed that
some plants were given signatures "in order to
set man on the right track," but others were
left blank to encourage humans to discover
them.
Della Porta provided the definitive DOS ex-
position of the 16th and 17th centuries (Clubb
1965). According to della Porta (1658 [orig.
1558]), "... Bugloss and Orchanet bear seeds
like a Viper's head, and these are good to heal
their venomous bitings. Likewise Stone—crop
and Saxifrage are good to break the stone in a
man's bladder." Another follower of Paracelsus,
Oswald Croll was equally strong in his espousal
of DOS: "In like manner, herbs magically by
their signature ... manifest their interiors, con-
cealed in the occult silence of Nature" (Crollius
1669).
History of DOS since the 1900s
Five publications cited in the 1947-1996
index to Economic Botany refer to DOS (Ka-
plan et al. 2001). Vickery (1981) and Touw
(1982) cited historical references. Browner
(1985) concluded that two plants were "appar-
ently selected" for infertility due to their resem-
blance to human genitalia. Carlson (1986) wrote,
"The closer the resemblance of the [ginseng]
root is to the human figure, the greater is it
thought of as having associative signature." Ha-
zlet (1986) averred, "The medieval notion that
plant morphology can suggest medicinal use of a
plant ... was manifest in several Cabecar medic-
inal plants." More recently Dafni and Lev (2002)
described DOS in modern Israel folk medicine.
"Heart—shaped" leaves of Melissa officinalis were
employed to treat heart weakness (Dafni and
Lev 2002).
Critiques of DOS
Dodoens (1583) was perhaps the first to dis-
miss DOS: "... it is so changeable and uncertain
that, it seems absolutely unworthy of acceptance"
(translation from Arber 1988 [orig. 1912]). Ray
(1717 [orig. 1691]) expressed disbelief in DOS.
Hahnemann (1825), the founder of homeopathy,
ridiculed the theory: "I shall spell the ordinary
medical school the humiliation of reminding it of
the folly of those ancient physicians who, deter-
mining the medicinal powers of crude drugs from
their signatures."
Thiselton—Dyer (1889), who reviewed DOS in
the 16th and 17th centuries, found DOS reference
in most medical works and noted that it was
"treated with a seriousness characteristic of the

2007] BENNETT: DOCTRINE OF SIGNATURES 249

backward state of medical science ...... Read
(1927) suggested that the influence of DOS di-
minished beginning in the 1700s but Arber (1988
[orig. 1912]) argued that it persisted into the 19th
century.
Modern herbals, pharmacopoeias, textbooks,
and peer—reviewed literature are nearly univer-
sal in condemning DOS. It is called fanciful,
far—fetched, premodern, prescientific, primitive,
unreliable, and unscientific (Table 2). Simpson
and Ogorzaly (2001) stated that the idea "seems
absurd now but received great acclaim when it
was proposed. Luckily, it was soon displaced by
less subjective and more secular methods of de-
termining a plant's medical efficacy." Tyler (1994)
ranked it among his 10 False Tenets of Para-
herbalism. Barford and Kvist (1996) concluded
that Ecuadorian plants "used according to the
Doctrine of Signatures" are less promising phar-
macological leads than other medicinal plants.
Balee (1994) succinctly summarized the current
view, noting that most economic botanists reject
DOS.
A few scholars are neutral in their descriptions
of DOS. The fear of mandrake may have been a
"primitive" result of DOS but it probably saved
many lives according to Schultes and Hofmann
(1980). Davis and Yost (1983) wrote, "Plants with
a strong odor might repel symptoms in a manner
reminiscent of the doctrine of signatures." In ref-
erence to willow bark's efficacy iri treating rheu-
matic pains, Kreig (1964) noted that, `At least
one of these quaint beliefs had a fragment of
truth in it."
Fewer scholars interpret DOS in a positive light.
In describing the discovery of a digitalis—like sub-
stance in a Mexican Magnolia species with cordate
leaves, Plotkin (2000) commented, "As ludicrous
as it sounds, the doctrine has yielded at least one
medicinal compound in wide use until recently."
James (n.d.) supported the utility of DOS as a
mnemonic device: "In a period where most of the
world was still largely illiterate, it is likely that the
Doctrine of Signatures was useful as a mnemonic
aid for the apprentice who was learning by obser-
vation and rote." The late Stephen Jay Gould
(2000) provided the most honest assessment of
DOS:
I question our usual dismissal of this older ap-
proach as absurd, mystical, or even prescientific (in
any more than a purely chronological sense) ...
But how can we blame our forebears for not know-
ing what later generations would discover? We
might as well despise ourselves because our grand-
children will, no doubt, understand the world in a
different way.

TABLE 2. CRITIQUES OF THE DOCTRINE OF SIGNATURES.

Recent Assessments Source

"Viewed from a scientific standpoint, their theory and diagnosis Mooney (1992 [orig. 1900])
are entirely wrong."
"pseudo-scientific" Arber (1988 [orig. 1912])
"The modern educated man is apt to be cynical in his attitude to Read (1927)
remedies based upon such unscientific dogmas."
"primitive imagination correlating shapes of organs and shapes of herbs Buchanan (1991 [orig. 1938])
by magical impulse" + "myth"
"makes as much sense as believing that walnuts are good for mental Tyler (1994)
illness because their kernels resemble the brain"
"no scientific basis to this concept" Levetin and McMahon (1999)
"far-fetched" Sumner (2000)
"seems absurd now" Simpson and Ogorzaly (2001)
"unreliable, unscientific and absolutely contrary to the homeopathic Saine (2002)
methodology"
"pre-modern knowledge" Panese (2003)
"totally debunked" Reed (2005a)
"magical similarity was re-exhumed in an empirical and intuitive Bellavite et al. (2005)
manner without any scientific understanding or experimental proof "
"fanciful medical theory" EncyclopEadia Britannica (2006b)
"pseudoscientific methods" EncyclopEadia Britannica (2006b)
250 ECONOMIC BOTANY [VOL. 61
Reinterpreting DOS
Both critics and proponents of DOS assume
that there is no diffusion of knowledge. Accord-
ing to their arguments, one would have to admit
that it is used for continual rediscovery of a
plant's medicinal value. In traditional cultures,
plant knowledge is effectively passed from one
generation to the next through observation and
oral tradition. DOS should be reevaluated with
respect to (1) its role in the discovery of medici-
nal plants, (2) post hoc attribution of signatures,
(3) the nature of signatures, and (4) its role as a
mnemonic.
DOS AND DISCOVERY
OF MEDICINAL PLANTS
Scholars were not present when healers in tra-
ditional cultures first learned of the value of me-
dicinal plants. Therefore, it is impossible to say
whether signatures influenced plant selection. If
DOS was a ubiquitous method of choosing me-
dicinal plants, as many argue, then plants bearing
signatures should be more widely used than those
lacking a signature. For example, one would ex-
pect many examples of heart—shaped leaves in
botanical pharmacopoeias for cardiac disease.
Is a plant's heart—shaped leaf a signature indi-
cating the plant's utility in treating cardiac condi-
tions? To address this question, I searched the
International Plant Name Index (http://www
.ipni.org) for six specific epithets that refer to
heart—shaped leaves. The search yielded 2,584 bi-
nomials. I then randomly selected 80 binomials
(ca. 3% of the total), sampling each epithet pro-
portional to its occurrence (Table 3). For invalid
names, I substituted another binomial with the
same specific epithet. Next, I searched the print
and electronic literature using both accepted
names and common synonyms for ethnobotanical
usage and pharmacological studies. Consulted lit-
erature included the Journal ofEthnopharmacology,
Schultes and Raffauf 's The Healing Forest: Medic-
inal and Toxic Plants of the Northwest Amazonia
(1990), Moerman's Native American Ethnobotany
(n.d.), Duke's Phytochemical and Ethnobotanical
Databases (n.d.), U.S. National Library of Medi-
cine's PubMed, and Prelude Medicinal Plants
Database for Africa (Baerts—Lehmann and
Lehmann n.d.).
Of the 80 species, 21 are used medicinally.
Three of the species are employed in a way that
may relate to cardiac medicine. Alchornea cordifolia
TABLE 3. OCCURRENCE OF EPITHETS RELATING TO
HEART-SHAPED LEAVES IN THE INTERNATIONAL
PLANT NAME INDEX.
Epithet Occurrences
cordata 980
cordatum 303
cordatus 115
cordifolia 824
cordifolium 243
cordifolius 119
Total 2,584
(Schumach. & Thonn. in C.F. Schumacher)
Mull. Arg., which has red fruits and cordate
leaves, is a smooth muscle relaxant and a diuretic
(Ogungbamila and Samuelsson 1990). Antigonon
leptopus Hook. & Arn, has antithrombin activity
(Chistokhodova et al. 2002). Sida cordifolia L.,
effective in treating brachycardia, is the only
species that clearly has cardiac value (Medeiros et
al. 2006). At most, 3 of the 80 species (or less
than 4%) are utilized in manner consistent with
DOS. These data clearly refute any a priori value
of heart—shaped leaves as signs for cardiac activity.
A second set of 80 randomly selected species,
without respect to specific epithets was analyzed
similarly. Of this data set, 17 species were em-
ployed in traditional medicine. The proportion
of medicinal plants in this sample and the
heart—shaped leaf sample do not differ statisti-
cally (Fishers Exact test, two—tailed probability
P=0.578). In other words, a plant with a distinct
sign is no more likely to be used than an arbitrar-
ily selected one.
POST HOC ATTRIBUTION
A second problem encountered when examin-
ing DOS is that signatures were applied after
plants had been incorporated into pharma-
copoeias. Read (1927) commented, ".. . in some
cases the signatures of drugs were observed after
their real use had been discovered." Reed (2005a)
would concur: "It is likely that many valuable
herbs were in use before the doctrine and that the
organ—plant match was made later to accommo-
date and validate the doctrine."
Consider the signature of willow bark cited by
Kreig (1964) and many others. In a review of
anti—inflammatory drugs, Vane and Botting
(1998) provided the following historical account
of its discovery.

2007] BENNETT: DOCTRINE OF SIGNATURES
On June 2, 1763, Edward Stone read a report to
the Royal Society on the use of willow bark in fever.
He had accidentally tasted it, and was surprised by its
extraordinary bitterness, which reminded him of the
taste of cinchona bark (containing quinine), then
being used to treat malaria. He believed in the 'doc-
trine of signatures', which dictated that cures for dis-
eases would be found in the same locations where the
malady occurs. Since the `willow delights in a moist
and wet soil, where agues chiefly abound', he gath-
ered a pound of willow bark, dried it over baker's
oven for 3 months, then ground it to a powder.
Rev. Stone "accidentally" tasted the bark. He
did not select willow because of a priori morpho-
logical cues. The parson may have been motivated
by the willow's growth in wet soil and the belief
that "remedies lie not far from their causes."
However, many other species occupy the same en-
vironment. Stone was objective in his approach.
"I have no other motives for publishing this valu-
able specific, than it may have a fair and full trial
in all its variety and circumstances and situations"
(in Vane and Botting 1998). Morphological cues
are insufficient to explain his selection.
Mooney (1992 [orig. 1900]) denigrates tradi-
tional Cherokee medicines writing, "... the doc-
tor puts into the decoction intended as a ver-
mifuge some of the red fleshy stalks of the
common purslane ... because these stalks some-
what resemble worms and consequently must
have some occult influence over worms." Dafni
and Lev (2002) also reveal the potential for pre-
dispositions to influence findings: "When the an-
swer was suspected by us to be related to the
Doctrine of Signatures, the next questions was
`Why this plant is used for this specific pur-
pose?"' Hazlett (1986), in reference to the red
spots on Columnea crassa leaves, says, "This `sign'
was interpreted as an indication that the plant is
effective in stopping nosebleeds." Here, the pas-
sive voice makes it unclear as to who interpreted
the signature. Browner (1985) asserted that two
species are "are apparently selected" based on
DOS. Saine (2002) criticized second–hand as-
sessments, noting "At best, these post hoc associa-
tions make studying the materia medica quaint
and colorful, but should never be confused with a
priori postulations used for prescribing accord-
ingly to the doctrine of signatures."
NATURE OF SIGNATURES
The application of signatures is not only post
hoc, but also elastic and inconsistent (Arber 1988
251
[orig. 1912]). A signature, like beauty, is in the eye
of the beholder. In 1628, Guy de la Brosse noted
that it was easy to imagine any 'resemblance be-
tween a plant and an animal that happened to be
convenient (cited in Arber 1988 [orig. 1912]). Ray
(1717 [orig. 1691]) considered signatures to be
"rather fancied by men than designed by Nature."
Buchanan (1991 [orig. 1938]) cited foxglove's
heart–shaped leaves, as a classic example of
DOS. However, the leaves of foxglove are not
heart–shaped. Cardiac glycosides are obtained
from two species of foxglove. One has lanceolate
leaves; foliage of the other is broadly ovate (Bailey
et al. 1976). It is as difficult to find a heart signa-
ture in Digitalis leaves as it is to visualize human
sex organs in the examples cited by Browner
(1985).
A second way to reevaluate DOS is to recon-
sider the nature of signatures. The coiled roots of
snakeroot for snakebite (Levetin and McMahon
1999), the clinging fruits of wild comfrey for
memory (Reed 2005b), and the eye–like seed of
aconite for ocular problem (Noth 1998) fit the
classical morphological definition. Signatures,
however, need not be morphological. A common
feature of medicinal pharmacopoeias is a large
number of plants with strong odors (e.g. Ankli et
al. 1999; Leonti et al. 2002; Shepard 2002). If
there were laws of ethnobotany, one would be
this: Plants with potent odors and strong tastes are
employed in medicine and ritual. These sensory
cues are correlated with the presence of bioactive
compounds. It is no coincidence that herbs and
spices possess antimicrobial activity (e.g., Billing
and Sherman 1998; Sherman and Billing 1999).
Expanding signatures to include olfactory or gus-
tatory clues is entirely sensible. The human nose
functions in the field the same way that a gas
chromatograph functions in the lab—both are
effective in detecting volatile compounds.
Debus (1973) noted, "The chemist's [Paracel-
sus's] own laboratory procedures seemed to offer
the proper key to nature's hidden secrets." Davis
and Yost's (1983) suggestion that plants with a
strong odor might repel symptoms echoes a simi-
lar view. Etkin (1988) argued that the selection of
some red plant makes sense due to the presence
of antimicrobial and haemostatic red quinones.
ROLE AS A MNEMONIC
Whether DOS ever led to the discovery of me-
dicinal plants is not testable. Leonti et al. (2002)
acknowledged the epistemological problems of
252 ECONOMIC BOTANY [VOL. 61
assigning an elucidation role to the organoleptic
properties of medicines. Moreover, many plants
that lack signatures are used for the same purpose
as those that have signatures. The Shuar employ at
least nine species to stanch bleeding (Bennett at
al. 2002). With its red—splotched leaves, machan-
cha is the only one of them that bears a sign that
could be interpreted as blood.
Many species with obvious "signatures" are not
employed medicinally. Association with a signa-
ture makes it easier to remember a plant and to
transmit knowledge about its use. One would ex-
pect signatures to converge with plants that truly
are efficacious. The Gnobe in Panama use the
bright red seeds of soroci as a blood tonic (Ben-
nett, unpub. field notes from June 1999, July
2000, June 2001), as do healers on Andros in the
Bahamas (Bennett, unpub. field notes from
March 2003). Chaturvedi (2005) demonstrated
that extracts from the plant normalized blood
glucose level, reduced triglyceride and LDL lev-
els, and increased HDL level. Other species bear-
ing red fruits or seeds are not included in the
pharmacopoeias.
Though it is unlikely that signatures led to dis-
covery of their medicinal values, many plants that
bear signatures are efficacious. Mooney's (1992
[orig. 1900]) dismissal of purslane was prema-
ture. The plant is effective in controlling intes-
tinal parasite loads and has gastroprotective
activity (Karimi et al. 2004; Quinlan et al. 2002).
Resemblance to worms aided in transmission of
knowledge about the plant's use. Eyebright eye
drops effectively and safely treat conjunctivitis
(Stoss et al. 2000). Eyebright was likely used for
ocular ailments because it worked and its signa-
ture facilitated transmission of the knowledge of
its use.
DOS utility as a memory aid was first prof-
fered in 1938 by Buchanan (1991 [orig. 1938])
and later mentioned by Etkin (1988), James
(n.d.), and Leonti et al. (2002). In his discussion
of the transmission of medicinal plant knowledge
by the Matisgenka and Yora of Peru, Shepard
(2002) supported this role: "In these and other
orally transmitted systems of thought, mnemonic
cues may be essential to the viability of knowl-
edge transmission. Plants that are both efficacious
and easy to remember are more likely to be main-
tained in the pharmacopoeia of non—literate soci-
eties through time."
The role of mnemonic in cultural transmission
is well established. Physicians, burdened with vast
indications, counter indications, and cures use a
simple mnemonic to remind them of anticholin-
ergic poisonings: red as a beet, hot as a hare,
blind as a bat, dry as a bone, and mad as a hatter.
Berlin (1992) found that "... many folk generic
names are metaphorically descriptive of some fea-
tures of the plant or animal species to which they
are applied, this to serving as a useful semantic
aid to memory." In his discussion of memory and
culture, Donald (1997) described "various forms
of visual imagination as a means of understand-
ing and retaining quite complex memories."
While not explicit in his argument, plant signa-
tures would seemingly qualify as visual imagina-
tions.
My fieldwork with the highland Quechua of
Peru, the Shuar and Quichua of Amazonian Ec-
uador, the Chachi of coastal Ecuador, the Gnobe
of Panama, and the Seminoles of southern
Florida has revealed many examples of plants
with apparent signatures. A Quichua healer, Luis
Cerda, showed me a red—tipped puca panga leaf.
The plant was employed to treat menstrual bleed-
ing and used for the treatment of snakebites. It
would have been tempting to ascribe the use of
this gesneriad to the DOS, but Luis never said
that it was used because the leaf was red. Several
years later, I collected a congeneric species on the
opposite side of the Andes. Horacio Lopez, a
Chachi Shaman provided the same uses, but his
gesneriad lacked the red signature. The Shuar,
with a pharmacopoeia exceeding 200 species,
treat liver ailments with a decoction made from
liver—shaped leaves of akapmas (Bennett et al.
2002). The association between the leaf's shape
and its use is a way of remembering and trans-
mitting knowledge of the plant's utility.
Buchanan (1991 [orig. 1938]) wrote, `Actually
it [DOS] is the mnemonic distillate of what must
have been considerable experience." The doctrine
of signatures is primarily a way of remembering
and transmitting plant knowledge, not a means
of discovery. Shepard (2002) strongly defends
DOS when he writes, "Western scientists should
reflect on their own habits before dismissing such
mnemonic devices as irrational or pre—scientific."
DOS is a meaningful device useful in disseminat-
ing knowledge of medicinal plants.
Conclusions
The Doctrine of Signatures is found throughout
the world. It was cited by the classical Greek writ-
ers and was used frequently in herbal medicine

2007] BENNETT: DOCTRINE OF SIGNATURES
during the Renaissance. With few exceptions,
scholars dismiss DOS as a "primitive" or "presci-
entific" idea. Despite its long history, the doctrine
has had little critical review. A careful evaluation
of signatures suggests four things. (1) As classi-
cally defined by morphology, signatures seldom if
ever led to the discovery of medicinal plants.
Considering DOS in this manner is unproduc-
tive and largely untestable. (2) Most signatures
are post hoc appellations rather than a priori
clues. Seeing a particular signature often requires
a vivid imagination. (3) A broader concept of sig-
natures that includes organoleptic properties as-
sociated with therapeutic value is productive.
Plants with strong odors and bitter tastes, even
those with no history of medicinal use, warrant
pharmacological investigation as these properties
are associated with the presence of potentially
bioactive compounds. (4) DOS should be consid-
ered for what it primarily is-a way of disseminat-
ing information. Fundamentally, the DOS is a
mnemonic and therefore it is exceedingly valuable
in traditional cultures. A fifth point warrants in-
vestigation. Moerman (2002, and pers. comm.)
suggests that signatures may enhance the "placebo
effect" of medicines. Those species with character-
istics resembling the disease being treated may be
perceived to be more effective by patients.
Acknowledgments
I thank Paul Cox for encouraging me to de-
velop this topic for publication and Daniel Moer-
man for his insightful comments on earlier man-
uscript drafts. This paper is contribution 128 in
Florida International University's "Tropical Biol-
ogy Publication" series.
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